MCB 32 Fall 2016

Lecture 14 (Oct 13)

Lecture 14 Skeletal muscle contraction

Warm-up activity
Remember that there are three types of muscle tissue (Fig 12.33):
1) Skeletal muscle attached to bones, control body movement, striated
Controlled voluntarily by somatic peripheral nervous system (motor neurons)
2) Cardiac muscle muscle in heart, striated
Controlled by autonomic NS and hormones (involuntary control)
3) Smooth muscle muscle in organs and tubes, moves material through body, smooth
Controlled by autonomic NS and hormones (involuntary control)
Today we will learn about skeletal muscle. You will learn more about the function and
regulation of cardiac and smooth muscle later in the course.
Skeletal muscle cell structure
Muscles attach to bones by tendons made of collagen (connective tissue)
Muscle is made of muscle fibers, blood vessels, nerves, connective tissue (Fig 12.1)
Each muscle fiber is a muscle cell that contains more than one nucleus. The fibers run the
entire length of the muscle.
Muscle fibers have some specialized cell structures (Fig 12.2):
Sarcolemma (membrane), sarcoplasm (cytoplasm), sarcoplasmic reticulum (ER)
Myofibril made of actin+myosin = what actually causes muscle contraction (Fig 12.3)
T-tubules long tubes in the sarcolemma, allows AP to get into middle of muscle fiber
Myofibrils create the force during muscle contraction. They are made up of two fibers:
1) Thick fiber made of myosin = motor protein, head region is flexible and moveable
and acts like a hinge, can bind actin, gets energy from ATP (Fig 12.5a)

2) Thin fiber made of actin = two F-actin polymers twisted around each other
Tropomyosin protein wraps around actin and troponin binds tropomyosin (Fig 12.4b)

MCB 32 Fall 2016

Lecture 14 (Oct 13)

Myosin heads bind actin, forming crossbridges. Myosin can walk along actin, moving the
actin filaments (slide)
The fibers are found in regular, repeating arrangements called sarcomeres. Sarcomeres are
what make the muscle look striated (Fig 12.5)
The Z lines are the ends of each sarcomere
Video of muscle contraction
Clicker question
How muscles generate force
During a muscle contraction, the myosin heads walk along actin filaments, bringing the ends
of the sarcomere (Z lines) closer together. This is known as the sliding filament model.
Fig 12.6
Note: The myosin and actin fibers are not changing length, they are just sliding past each other,
which shortens the sarcomere.
How does myosin walk along actin?
This is called the crossbridge cycle and requires ATP hydrolysis. Basically, the myosin head
will bind actin, then the head will pivot (the power stroke), moving the actin with it. After
binding an ATP, the myosin will then release actin and reattach further down the actin filament
for another stroke. (Fig 12.7 + animation)
Why dont filaments slide all the way back to their original position each time a myosin head
releases?

Why does death cause rigor mortis?

Excitation-contraction coupling
Clicker question review
Now we know how myosin and actin are able to generate force in muscles, but we also know
that muscle contraction only happens when a motor neuron releases ACh onto the muscle.

MCB 32 Fall 2016

Lecture 14 (Oct 13)

How does excitation of the muscle (via ACh binding receptors) couple to muscle contraction?
1) Motor neuron releases ACh onto muscle
2) nAChR open, let in Na+ which depolarizes muscle (always excitatory)
3) Voltage gated Na+ channels open in muscle muscle AP (same as neuron AP)
Release of ACh onto muscle always triggers AP (unlike neuronal graded potential)
4) AP travels down T-tubule into interior of muscle
5) AP causes calcium (Ca+2) release channels to open in sarcoplasmic reticulum (SR), which
stores high concentrations of Ca+2 (calcium goes down concentration gradient into cytoplasm)
6) Calcium from SR floods cytoplasm, binds troponin and exposes myosin binding site on actin
7) Myosin walks down actin filaments, decreasing length of sarcomere (Fig 12.8)

How does an increase in intracellular Ca+2 lead to myosin binding?

- Tropomyosin blocks the binding sites for myosin on actin in low Ca+2. Myosin can bind, but
its a weak interaction.
- When calcium levels in cytoplasm increase, calcium binds troponin, which pulls tropomyosin
over, exposing binding sites for myosin heads (Fig 12.9). Myosin cannot properly move along
actin until tropomyosin moves out of the way.
3

MCB 32 Fall 2016

Lecture 14 (Oct 13)

When the motor neuron stops firing action potentials, ACh will be degraded by AChE and the
muscle will return to its resting state by pumping Ca+2 back into the SR and tropomyosin will
cover actin again, so myosin cant bind.
*Summary animation posted on bCourses
Clicker question
Muscle metabolism
Muscle contraction requires a lot of ATP (why?) and it must be readily available.
Review of aerobic vs anaerobic respiration (Fig 12.22)
Muscles have a small amount of ATP ready at rest, but they also store energy in the bonds of a
molecule called creatine phosphate.

When a muscle cell is working, ADP levels rise, which drives the reaction to the right, making
ATP
During exercise, the energy from creatine phosphate will last for less than a minute, but this is
enough time for cellular respiration to increase ATP production.
At first the muscles use their own stores of glycogen to supply glucose for cellular respiration,
but after those stores are used up, they get glucose and fatty acids delivered by the blood
During strenuous exercise, there isnt enough oxygen to support oxidative phosphorylation, so
the muscles must rely on anaerobic glycolysis as well. (Fig 12.23)

Which athletes would benefit the most from creatine supplements? (slide)

MCB 32 Fall 2016

Lecture 14 (Oct 13)

From Twitch to Contraction

Video of real muscle contraction
Clicker
A twitch is when a muscle cell responds to a single action potential. Twitches are always the
same duration and produce the same amount of tension (all or nothing).
We know that muscles can produce different amounts of force. How is that possible if a single
twitch is always the same?
1) Summation in a single muscle fiber (also known as temporal summation)
If a muscle fiber is stimulated at higher and higher frequencies, the resulting tension can
summate beyond the amplitude of a single twitch
If stimulation frequency is fast enough, the muscle wont get a chance to relax and Ca+2 wont
be pumped back into SR before the next stimulation. This can lead to fused tetanus muscle
doesnt relax at all between stimuli, so you have a sustained maximal contraction. (Fig 12.16)
Most conscious muscle movement utilizes tetanus
2) Recruitment of multiple motor units (also known as spatial summation)
Remember that one motor neuron can innervate multiple muscle fibers. When the motor
neuron fires an action potential, all of the fibers in the motor unit will contract.
The more motor units within a muscle that are recruited into a contraction, the stronger the
force generated by the muscle (Fig 12.18)
Motor units are usually recruited by size, so smaller motor units are used first and then bigger
and bigger motor units get recruited as the muscle force increases. (Fig 12.19)
Fiber length vs tension
Another way to get maximal tension is by making sure the sarcomeres are at an optimal,
intermediate length (Fig 12.17)
Too long: myosin-actin form just a few crossbridges
Too short: only a little bit of distance left to move thin filaments
In lecture 15, we will learn about the different types of muscle fibers and how they are
controlled by the CNS